Competing theories of prefrontal Cortex Functions:  Processes, representations and gaps

Jordan Grafman
NINDS
Bethesda, MD


Course Prerequisites

Introductory graduate level coursework in cognitive psychology, cognitive neuroscience, and neuroscience.


 Course description
  

The prefrontal cortex (PFC) has been implicated in a variety of “higher” cognitive functions – language, abstract reasoning, problem solving, social interactions, and planning. A number of theories have been proposed for how the PFC might mediate these functions.
This course will focus on strategies you can use to think about how to characterize the functions of the human prefrontal cortex.  I will briefly discuss the evolution, development, and neuroanatomy of the human prefrontal cortex followed by a summary of the advantages and pitfalls of studying normal volunteers and patients using both functional neuroimaging and the lesion approach.  Then, I will present the five criteria that a theory should meet if it is to provide a useful framework for understanding the functions of the prefrontal cortex. The bulk of the course will be an overview of some of the key PFC theories and my determination of how well they meet this criteria.  The criteria by which the theories will be assessed are: (1) specification of the type of information stored in the prefrontal cortex; (2) consistency with our knowledge of stimulus representation in the brain; (3) consistency with what is known of the evolutionary development of the prefrontal cortex; (4) ability to test the model and hence to verify or invalidate it; and (5) consistency of the model with available experimental data.  A theory may take a processing approach – i.e., specify computational procedures that are performed by the prefrontal cortex in order to manipulate information stored elsewhere in the brain. Alternatively, a theory may take a representational approach – i.e., specify the type of information stored in memories in the PFC. Finally, a theory might have components of both the processing and representational viewpoints.
I will discuss the main claims of each key theory and review data addressing these claims. You will learn that the models meet the above criteria to varying degrees. While all models are supported to some extent by the available cognitive neuroscience data, not all of the models address all of the available data. In particular, many researchers rely almost solely on functional neuroimaging data ignoring other sources of evidence (e.g., lesion studies).  With respect to specific theories, without modification, no single theory of PFC function appears to explain all of the available data.
Finally, I will argue that the representational approach seems to be more consistent with the above criteria than does the processing approach.  I will show how the representational approach forces a more detailed specification of a theory and thus enables specific hypothesis testing and is the most parsimonious way to explore the nature of knowledge stored in the human PFC.



Reading List

Required Readings

•    Wood, J.N. and Grafman, J.  Human Prefrontal Cortex: processing and representational perspectives. Nature Reviews Neuroscience, 4 (2), 139-147, 2003.

•    Moll, J., de Oliveira-Souza, R., Bramati I.E., and Grafman, J. Functional networks in emotional moral and non-moral social judgments.  Neuroimage,16 (3), 696-703, 2002.

•    Zalla, T., Phipps, M., and Grafman, J.  Story Processing in Patients with Damage to the Prefrontal Cortex.  Cortex, 38 (2), 215-231, 2002.

•    Rattermann, M.J., Spector, L., Grafman, J., Levin, H., and Howard, H.  Partial and Total-Order Planning: Evidence from Normal and Prefrontally Damaged Populations.  Cognitive Science,25 (6), 941-975, 2001.

•    Milne, E., and Grafman, J.  Ventromedial Prefrontal Cortex Lesions in Humans Eliminate Implicit Gender Stereotyping. Journal of Neuroscience, 21 (RC150), 1-6, 2001.

•    Grafman, J., and Litvan, I.  Recognizing the importance of deficits in executive functions.  Lancet, 354 (9194/4 December), 1921-1923, 1999.


Optional Readings


•    Grafman, J. Experimental Assessment of Adult Frontal Lobe Function.  In: B.L. Miller and J. Cummings (Eds.) The Human Frontal Lobes: Function and Disorder. New York: The Guilford Press,321-344, 1999.

•    Grafman, J. The Structured Event Complex and the Human Prefrontal Cortex.  In: D.T. Stuss and R.T. Knight (Eds.) The Frontal Lobes. New York: Oxford University Press, 2002.

•    Koechlin, E., Danek, A., Burnod, Y., and Grafman, J.  Medial Prefrontal and Subcortical Mechanisms Underlying the Acquisition  of Motor and Cognitive Action Sequences in Humans.  Neuron, 35 (2), 371-381, 2002.

•    Wood, J.N., Romero, S.G., Makale, M., & Grafman, J.  Category-specific representations of social and non-social knowledge in the human prefrontal cortex.  Journal of Cognitive Neuroscience, 15 (2), 236-248, 2003.

•    Dreher, J.C., and Grafman, J.  Dissociating the Roles of the Rostral Anterior Cingulate and the Lateral Prefrontal Cortices in Performing Two Tasks Simultaneously or Successively.  Cerebral Cortex, 13 (4), 329-339, 2003.

•    Rosen, V.M., Caplan, L., Sheesley, L., Rodriquez, R., and Grafman, J.  An examination of daily activities and their scripts across the adult lifespan.  Behavior Research Methods, Instruments, and Computers,35 (1), 32-48, 2003.

•    Sanfey, A.G., Hastie, R., Colvin, M.K., and Grafman, J.  Phineas Gauged: Decision-Making and the Human Prefrontal Cortex.  Neuropsychologia, 41 (9), 1218-1229, 2003.

•    Fuster, J. The Prefrontal Cortex (3rd Edition). New York: Raven Press, 1997.

•    Macmillan, M. An Odd Kind of Fame: Stories of Phineas Gage.  Cambridge: MIT Press, 2000.

•    Miller, E.K. and Cohen, J.D.  An Integrative Theory of Prefrontal Cortex Function.  Annual Review of Neuroscience, 24, 167-202, 2001.



Course Presentation

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